Louisiana State University LSU Digital Commons LSU Historical Dissertations and Theses Graduate School 1965 The rC anial Osteology of the New World Tyrannoidea and Its Taxonomic Implications. Stuart Lawrence Warter Louisiana State University and Agricultural & Mechanical College Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_disstheses Recommended Citation Warter, Stuart Lawrence, "The rC anial Osteology of the New World Tyrannoidea and Its Taxonomic Implications." (1965). LSU Historical Dissertations and Theses. 1105. https://digitalcommons.lsu.edu/gradschool_disstheses/1105 This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Historical Dissertations and Theses by an authorized administrator of LSU Digital Commons. For more information, please contact [email protected]. This dissertation has been microfilmed exactly as received WAKTPR, Stuart lawrciH'c*, lf)M4— TUP CRANIAL OSTPOIAXiY OF TH K NKW WORLD TYRANNOIJ )KA ANU ITS TAXONOMIC IMPLICATIONS. I xjuisiaua State University, Ph.D., 1005 / . o o l o g y University Microiihns, Inc,, Ann Arbor, Michigan THE CRANIAL OSTEOLOGY OF THE NEW WORLD TYRANNOIDEA AND ITS TAXONOMIC IMPLICATIONS A Dissertation Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy in The Department of Zoology and Physiology by Stuart Lawrence Warter B.S., University of Miami, 1956 M.S., University of Miami, 1958 August, 1965 ACKNOWLEDGMENTS Appreciation 10 expE«*ia*d to the many individuals end organizations that have generously provided assistance in the pursuit and completion of this study. The following individuals graciously allowed examina­ tion of materials in collections in their care: Dr. Pierce Brodkorb, University of Florida, Gainesville; Dr. Philip S. Htanphrey, United States National Museum, Washington; Dr. Richard F. Johnston, University of Kansas, Lawrence; Dr. Raymond Paynter, Harvard University, Cambridge; Dr. Kenneth C. Parkes, Carnegie Museum, Pittsburgh; Dr. Robert W. Storer, University of Michigan, Ann Arbor. Dr. Dean Amadon, American Museum of Natural History, New York, and Drs. Humphrey and Storer generously provided the loan of specimens for protracted study. Dr. Peter L. Ames of the University of California, Berkeley, Dr* Mary A. Heimerdinger of the Carnegie Museum, Dr. Richard L. Zusi of the U. S. National Museum, and numerous other ornithologists provided advice, information, encouragement, and enlightening discussions that contrib­ uted significantly to the ideas and conclusions presented in this work. The Department of Zoology, Louisiana State University, il provided travel funds for one of the several trips a>ade to museuas for the examination of material. Marcia Brady Tucker and the American Omithologlsts' Union provided expenses that enabled the presentation of some of the results of this study before the above organi­ zation in convention at Lawrence, Kansas, in August, 1964. Dr. Murray S. Blum provided support and encouragement during the period of our association in the Department of Entomology. Dr. Robert J. Newman of the Museum of Zoology kindly donated his time in the critical reading of this manuscript. I am particularly indebted to Dr. George H. Lowery, Jr., for his enthusiastic and unfailing support and guidance throughout the course of his direction of this study. ill TABLE OF CONTENTS PAGE TITLE P A G E ...... 1 ACKNOWLEDGMENTS .................................... It TABLE OF CONTENTS .......................... tv LIST OF TABLES .................................... vt LIST OF FIGURES ............................. ........vti ABSTRACT ...........................................vitl INTRODUCTION ...................................... 1 CLASSIFICATION OF THE TYRANNOIDEA .................. 12 DESCRIPTIONS ...... 19 Introduction .................................... 19 General Description .............................. 19 Type I Skulls ................ .............. ..... 26 Type II Skulls ................................ 39 Type III Skulls .................................. 47 Type IV Skulls ... 51 Type V Skulls .................................... 51 Type VI Skulls ................................... 55 PROMISING TAXONOMIC CHARACTERS ..................... 56 The Nasal Region .......................... 56 The Palatine Process of the Premaxllla (Palato­ maxillary ...... 73 iv PAGE DISCUSSION ........................................ 85 General Considerations ........................... 85 The Tyrannidae and Unsolved Problems ............. 93 Relationships of the Tityras and Becards ......... 97 The Status of the Pipridae ....................... 101 The Larger Fruit-eaters ........................... 117 TAXONOMIC CONCLUSIONS .................. ........... 128 Introduction ..................................... 128 Family TYRANNIDAE ......................... 130 Family PIPRIDAE ................................. 131 Family COTINGIDAE ................................ 133 Family RUPICOLIDAE ............................... 141 Family PHYTOTOMIDAE .............................. 142 Family OXYRUNC IDAS ............................... 143 SUMMARY AND CONCLUSIONS ............................ 145 LITERATURE CITED ................................... 148 APPENDIX I ........................................ 155 APPENDIX II ....................................... 159 VITA ............................................... 161 LIST OF TABLES TABLE PAGE I. Generalized Correlations of Skull Types with Feeding Habits and I'xternal Features ..... 18 II. Variable Characters or Flycatcher Skulls .... 31 III. Types of Nasal Septa in Representative Genera .................................. 70 vi LIST OF FIGURES FIGUPJE PAGE 1. Cranial Features of Tvrannua dominieana la .... 22 2. Types of Nasal Septa ........................ 28 3. Features of Type I Skulls ............ 30 4. Type II and III Skulls ...................... 41 3. Type V and VI Skulls ........................ 53 6. The Tyrannoid Nasal Region ................... 59 7. Principal Variants of the Typical Tyrannoid Nasal Capsule ............................ 63 8. Reconstructions of Tyrannoid Nasal Capsules ... 67 9. Representative Nasal Trabeculae .......... 72 10. Palatine Process of the Premaxilla ........... 78 11. Possible Derivation of Some Tyrannoidea Based on Type I and III Skulls ............. 103 12. Hypothetical Derivation of Exaspidean Tarsal Scutellation Among the Tyrannoidea ........ 109 13. Cranial Features of Sapavoa and Representative Kurylaimld Genera ........................ 115 14. Skull of Procnlas nudlcollls ................. 119 vii ABSTRACT The six families of Neotropical tyrannoid passerine birds, comprised of over 500 species In approximately 180 genera, constitute a major portion of the avifauna of South and Central America. The last major revision of this group was completed more than 35 years ago. In an effort to provide at least partial solutions to some of the existing systematic problems, a survey of the cranial osteology of members of these families was under­ taken. Nearly 1200 skulls of 224 species, comprising 117 genera, were examined. The skulls examined were placed within morphological groupings that seem to correspond closely to groupings based on other known biological features of the birds. Characters of the feet and legs had been relied upon previously to place within families the birds for which the more basic anatomical features were not known. Some of these placements have seemed incorrect to many ornitholo­ gists but no other information was available for evaluation of these allocations. The skull, in combination with general appearance, food habits, and breeding behavior (insofar as is known), seems to provide a character complex that helps to clarify relationships among this difficult viii group of birds. Skulls of Che flycatchers (Tyrannidae) are relatively uniform when compared to those of the Cotingidae. Several types of skulls occur within the diverse family Cotingidae; these types correspond to some of the subfamily groupings which, although widely used in the last century, have now disappeared from the literature. The use of several of these subfamilles--Attilinae, Tityrinae, Cotlnginae, Gymnoderlnae, and Querulinae--is reinstltuted. A new sub­ family is erected to contain the bellblrds (Procnlas). Skulls of most of the manakins (Pipridae) are barely distinct from those of the Cotlnginae, but the retention of the family is suggested since most of its members possess an elaborate method of courtship not yet found so well developed among the Cotingidae. The skulls of birds of the monotypic families Rupl- colidae, Phytotomidae, and Oxyruncldae were all found to be highly specialized and provide no evidence for uniting any of these birds with any other family. Skulls of the cotlngaa frtSJULft» > LL*£Xr ttnKtjptflM, Bra tor. and, to a lesser extent, TitYM flycatcher like. The mourners (Rhvt interna) are transferred to the Tyrannidae, and the reallocation of the other genera Is discussed. ix INTRODUCTION Although birds arc probably the most thoroughly studied class of vertebrates, the foundation on which avian classification rests in many cases is not substan­ tial. The ornithologist is only too aware of the inade­ quacies that exist at all levels of the present system, particularly with regard to the taxa above the genus. For the